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1. How much should an individual invest in reproduction as it grows older? Answering this question involves determining whether individuals measure their age as the time left for future reproduction or as the rate of deterioration in their state. Theory suggests that in the former case individuals should increase their allocation of resources to reproduction as opportunities for future breeding dwindle, and terminally invest when they breed for the last time. In the latter case they should reduce their investment in reproduction with age, either through adaptive reproductive restraint or as a passive by-product of senescence. 2. Here we present the results of experiments on female burying beetles, Nicrophorus vespilloides, in which we independently manipulated the perceived risk of death (by activating the immune system) and the extent of deterioration in state (by changing age of first reproduction and/or prior investment in reproduction). 3. We found that the risk of death and state each independently influenced the extent of reproductive investment. Specifically, we found a state-dependent decline in reproductive investment as females grew older that could be attributed to both adaptive reproductive restraint and senescence. A perceived increase in the risk of death, induced by activation of the immune system, caused females to switch from a strategy of reproductive restraint to terminal investment. Nevertheless, absolute reproductive investment was lower in older females, indicating constraints of senescence. 4. Our results show that a decline in reproductive investment with age does not necessarily constitute evidence of reproductive senescence but can also result from adaptive reproductive restraint. 5. Our results further suggest that the extent of reproductive investment is dependent on several different intrinsic cues and that the particular blend of cues available at any given age can yield very different patterns of investment. Perhaps this explains why a

Life history theory suggests that offspring desertion can be an adaptive reproductive strategy, in which parents forgo the costly care of an unprofitable current brood to save resources for future reproduction. In the burying beetle, Nicrophorus vespilloides, parents commonly abandon their offspring to the care of others, resulting in female-only care, male-only care, brood parasitism and the care of offspring sired by satellite males. Furthermore, when there is biparental care, males routinely desert the brood before larval development is complete, leaving females behind to tend their young. We attempted to understand these patterns of offspring desertion by using laboratory experiments to compare the fitness costs associated with parental care for each sex and the residual reproductive value of the two sexes. We also tested whether current brood size and residual reproductive value together predicted the incidence of brood desertion. We found that males and females each sustained fecundity costs as a consequence of caring for larvae, and that these costs were of comparable magnitude. Nevertheless, males had greater residual reproductive value than females and were more likely than females to desert experimental broods. Our results can explain why males desert the brood earlier than females in nature, and why female-only care is more common than male-only care. They also suggest that the tipping point from brood parasitism or satellite male behavior to communal breeding (and vice versa) depends on the value of the current brood relative to residual reproductive value.

1. A key component of parental care involves defending resources destined for offspring from a diverse array of potential interspecific competitors, such as social parasites, fungi and bacteria. 2. Just as with other aspects of parental care, such as offspring provisioning or brood defence, sexual conflict between parents may arise over how to share the costs of this form of care. There has been little previous work, however, to investigate how this particular burden might be shared. 3. Here we describe a hitherto uncharacterized form of parental care in burying beetles Nicrophorus vespilloides, a species which prepares carrion for its young and faces competition from microbes for this resource. We found that parents defend the carcass with antibacterial anal exudates, and that the antibacterial activity of these exudates is only upregulated following the discovery of a corpse. At the same time, phenoloxidase activity in the anal exudates is downregulated, indicating parallels with the internal insect immune system. 4. In unmanipulated breeding pairs, females had higher antibacterial activity in their anal exudates than males, suggesting sex-specific roles in this aspect of parental care. 5. When we experimentally widowed males, we found that they increased levels of antibacterial activity in their anal exudates. Experimentally widowing females, however, led them to decrease levels of antibacterial activity in their anal exudates. Widowed beetles of each sex thus produced anal exudates of comparable antibacterial activity. We suggest that this flexible division of antibacterial activity may be coordinated by Juvenile Hormone.

Parasites and pathogens are a ubiquitous threat facing all organisms. Life-history theory predicts that if investment in parasite resistance mechanisms is costly (as suggested by numerous studies), then organisms should tailor their investment in them to match their perceived risk of infection. Because most parasites are transmitted in a positively density-dependent manner, the threat from parasites tends to increase as population density increases. As a result, it is predicted that organisms should use population density as a cue to the risk of becoming infected and should increase investment in disease resistance mechanisms as the degree of crowding increases - this is known as density-dependent prophylaxis (DDP). This phenomenon has been experimentally tested in a number of insect species, and in most cases support for the DDP hypothesis has been forthcoming. DDP is likely to be particularly prevalent in species exhibiting density-dependent phase polyphenism (i.e. the phenotype adopted by the insect is plastic and dependent on the population density it experiences during its early development). We discuss the hormonal and genetic mechanisms underlying phase polyphenism and DDP, and speculate on the circumstances leading to their evolution. We end by discussing how future research into DDP might develop.

Maternal effects provide an important mechanism for mothers to create variation in offspring personality, and to potentially influence offspring life history strategies e.g. creating more/less dispersive phenotypes.However, within-clutch maternal effects often vary and hence there is potential for within-clutch varia-tion in personality. We studied the effects of hatching order on explorative and neophobic behaviour of the magpies Pica pica in relation to sex using novel environment and novel object experiments. Hatching order did affect explorative behaviour in magpie, but did so in opposite directions for either sex. First-hatched females were more explorative and had a tendency to be less neophobic, whereas in males, the reverse was true. Our results suggest that hormonal as well as post-natal environmental mechanisms could be underpinning this pattern. Future research is needed to fully understand the importance of both in creating different offspring personalities.

1. The risk of parasitism and infectious disease is expected to increase with population density as a consequence of positive density-dependent transmission rates. Therefore, species that encounter large fluctuations in population density are predicted to exhibit plasticity in their immune system, such that investment in costly immune defences is adjusted to match the probability of exposure to parasites and pathogens (i.e. density dependent prophylaxis). 2. Despite growing evidence that insects in high-density populations show the predicted increase in resistance to certain pathogens, few studies have examined the underlying alteration in immune function. As many of these species show increased cuticular melanism at high densities, the aim of this study was to use a multivariate approach to quantify relative variation in the allocation of resources to immunity associated with both rearing density (solitary vs. crowded) and cuticular colour (pale vs. dark) in a phase-polyphenic Lepidopteran species (Spodoptera littoralis Boisduval). 3. Relative to pale individuals, dark larvae (the high-density phenotype) exhibited higher haemolymph and cuticular phenoloxidase (PO) activity and a stronger melanotic encapsulation response to an artificial parasite inserted into the haemocoel. However, they also exhibited lower antibacterial (lysozyme-like) activity than pale larvae. Larval density per se had little effect on most of the immune parameters measured, though capsule melanization and antibacterial activity were significantly higher in solitary-reared than crowded larvae. 4. Correcting for variation in larval body condition, as estimated by weight and haemolymph protein levels, had little effect on these results, suggesting that variation in immune function across treatment groups cannot be explained by condition-dependence. These results are examined in relation to pathogen resistance, and the possibility of a trade-off within the immune system is discussed.

There is growing evidence that insects in high-density populations invest relatively more in pathogen resistance than those in low-density populations (i.e. density-dependent prophylaxis). Such increases in resistance are often accompanied by cuticular melanism, which is characteristic of the high-density form of many phase polyphenic insects. Both melanism and pathogen resistance involve the prophenoloxidase enzyme system. In this paper the link between resistance, melanism and phenoloxidase activity is examined in Spodoptera larvae. In S. exempta, cuticular melanism was positively correlated with phenoloxidase activity in the cuticle, haemolymph and midgut. Melanic S. exempta larvae were found to melanize a greater proportion of eggs of the ectoparasitoid Euplectrus laphygmae than non-melanic larvae, and melanic S. littoralis were more resistant to the entomopathogenic fungus Beauveria bassiana (in S. exempta the association between melanism and fungal resistance was non-sign®cant). These results strengthen the link between melanism and disease resistance and implicate the involvement of phenoloxidase.

Phenoloxidase (PO) is believed to be a key mediator of immune function in insects and has been implicated both in non-self recognition and in resistance to a variety of parasites and pathogens, including baculoviruses and parasitoids. Using larvae of the Egyptian cotton leafworm, Spodoptera littoralis, we found that despite its apparent importance, haemolymph PO activity varied markedly between individuals, even amongst insects reared under apparently identical conditions. Sib-analysis methods were used to determine whether individuals varied genetically in their PO activity, and hence in one aspect of immune function. Theheritability estimate of haemolymph PO activity was high (h2 = 0.690 ± 0.069), and PO activity in the haemolymph was strongly correlated with PO activity in both the cuticle and midgut; the sites of entry for most parasites and pathogens. Haemolymph PO activity was also strongly correlated with the degree to which a synthetic parasite (a small piece of nylon monofilament) was encapsulated and melanized (r = 0.622 ± 0.142), suggesting that the encapsulation response is also heritable. The mechanism maintaining this genetic variation has yet to be elucidated.

Theory predicts that natural selection will erode additive genetic variation in fitness-related traits. However, numerous studies have found considerable heritable variation in traits related to immune function, which should be closely linked to fitness. This could be due to trade-offs maintaining variation in these traits. We used the Egyptian cotton leafworm, Spodoptera littoralis, as a model system to examine the quantitative genetics of insect immune function. We estimated the heritabilities of several different measures of innate immunity and the genetic correlations between these immune traits and a number of life history traits. Our results provide the first evidence for a potential genetic trade-off within the insect immune system, with antibacterial activity (lysozyme-like) exhibiting a significant negative genetic correlation with haemocyte density, which itself is positively genetically correlated with both haemolymph phenoloxidase activity and cuticular melanization. We speculate on a potential trade-off between defence against parasites and predators, mediated by larval colour, and its role in maintaining genetic variation in traits under natural selection.

If a novel, resistant host plant genotype arises in the environment, insect populations utilising that host must be able to overcome that resistance in order that they can maintain their ability to feed on that host. The ability to evolve resistance to host-plant defences depends upon additive genetic variation in larval performance and adult host-choice preference. To investigate the potential of a generalist herbivore to respond to a novel resistant host we estimated the heritability of larval performance in the noctuid moth, Helicoverpa armigera, on a resistant and a susceptible variety of the chickpea, Cicer arietinum, at two different life-stages. Heritability estimates were higher for neonates than for third-instar larvae suggesting that their ability to establish on plants could be key to the evolution of resistance in this species, however further information regarding the nature of selection in the field would be required to confirm this prediction. There was no genetic correlation between larval performance and oviposition preference, indicating that female moths do not choose the most suitable plant for their offspring. We also found significant genotype by environment interactions for neonates (but not third-instar larvae), suggesting that the larval response to different plant genotypes is stage-specific in this species.

The eggs of amniotes exhibit a remarkable variety of shapes, from spherical to elongate and from symmetrical to asymmetrical. We examine eggshell geometry in a diverse sample of fossil and living amniotes using geometric morphometrics and linear measurements. Our goal is to quantify patterns of morphospace occupation and shape variation in the eggs of recent through to Mesozoic birds (neornithe plus non-neornithe avialans), as well as in eggs attributed to non-avialan theropods. In most amniotes, eggs show significant deviation from sphericity, but departure from symmetry around the equatorial axis is mostly confined to theropods and birds. Mesozoic bird eggs differ significantly from extant bird eggs, but extinct Cenozoic bird eggs do not. This suggests that the range of egg shapes in extant birds had already been attained in the Cenozoic. We conclude with a discussion of possible biological factors imparting variation to egg shapes during their formation in the oviduct.

Investment in immunity is costly, so that resource-based trade-offs between immunity and sexually-selected ornaments might be expected. The amount of resources that an individual can invest in each trait will be limited by the total resources available to them. It would therefore be informative to investigate how investment in immune function changes during growth or production of the sexual trait as resources are diverted to it. Using the dung beetle, Onthophagus taurus, which displays both sexual and male dimorphism in horn size, we examined changes in one measure of immune function, phenoloxidase activity, in the hemolymph of larvae prior to, and during horn growth. We found that phenoloxidase levels differed between small and large horned males throughout the final instar prior to the point where investment in horn growth was taking place. PO levels in females were intermediate to the two male morphs. These differences could not be accounted for by differences in condition, measured as hemolymph protein levels and weight. We suggest that the observed differences might be associated with sex and morph specific variation in juvenile hormone levels.

Several insect species show an increase in cuticular melanism in response to high densities. In some species, there is evidence that this melanism is correlated with an up-regulation of certain immune system components, particularly phenoloxidase (PO) activity, and with the down-regulation of lysozyme activity, suggesting a trade-off between the two traits. As melanism has a genetic component, we selected both melanic and nonmelanic lines of the phase-polyphenic lepidopteran, Spodoptera littoralis, in order to test for a causative genetic link between melanism, PO activity and lysozyme activity, and to establish if there are any life-history costs associated with the melanic response. We found that, in fact, melanic lines had lower PO activity and higher lysozyme activity than nonmelanic lines, confirming a genetic trade-off between the two immune responses, but also indicating a genetic trade-off between melanism and PO activity. In addition, we found that lines with high PO activity had slower development rates suggesting that investment in PO, rather than in melanism, is costly.

1. Mounting an immune response is likely to be costly in terms of energy and nutrients, and so it is predicted that dietary intake should change in response to infection to offset these costs. The present study focuses on the interactions between a specialist grass-feeding caterpillar species, the African armyworm Spodoptera exempta, and an opportunist bacterium, Bacillus subtilis. 2. The main aims of the study were (i) to establish the macronutrient costs to the insect host of surviving a systemic bacterial infection, (ii) to determine the relative importance of dietary protein and carbohydrate to immune system functions, and (iii) to determine whether there is an adaptive change in the host’s normal feeding behaviour in response to bacterial challenge, such that the nutritional costs of resisting infection are offset. 3. We show that the survival of bacterially-infected larvae increased with increasing dietary protein-to-carbohydrate (P:C) ratio, suggesting a protein cost associated with bacterial resistance. As dietary protein levels increased, there was an increase in antibacterial activity, phenoloxidase (PO) activity and protein levels in the haemolymph, providing a potential source for this protein cost. However, there was also evidence for a physiological trade-off between antibacterial activity and phenoloxidase activity, as larvae whose antibacterial activity levels were elevated in response to immune activation had reduced PO activity. 4. When given a choice between two diets varying in their P:C ratios, larvae injected with a sub-lethal dose of bacteria increased their protein intake relative to control larvae whilst maintaining similar carbohydrate intake levels. These results are consistent with the notion that S. exempta larvae alter their feeding behaviour in response to bacterial infection in a manner that is likely to enhance the levels of protein available for producing the immune system components required to resist bacterial infections (‘self-me

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It is well known that organisms defend their fitness against attack from parasites and pathogens by mounting a personal immune response. However, there is increasing evidence that organisms from diverse taxa also exhibit immune responses for the purpose of protecting other individuals as well as themselves. We argue that any type of immunity that has fitness consequences for both the challenged individual and one or more recipients should be referred to as ‘social immunity’. We show that social immune systems are a widespread yet relatively neglected component of immunity, ideal for the study of social evolution. Whereas personal immune systems protect lifespan, social immune systems effectively defend the fecundity component of fitness, commonly protecting offspring or reproductive kin. We suggest that there are likely to be close links between life history and the extent of investment in each form of immunity. Furthermore, trade-offs between social and personal immunity may explain individual variation in personal immune responses, including sex-specific immune defences.

Pigeons and humans are two highly visual species that have evolved separately for about 310 million years (Kumar and Hedges, 1998) and developed largely convergent visual systems due to similar visual needs. To investigatepigeon vision and cognitive abilities twodimensional pictorial stimuli are often used. However, it is not entirely clear, how pigeons perceive such stimuli and whether or not they can associate photographs with real objects. In the present study nine pigeons and eleven humans were trained to discriminate between photographs of two biologically irrelevant objects (“Greebles”). The pigeons were housed in an aviary containing the real Greebles and were trained in wooden chambers where they had to peck on a Plexiglas disk when positive stimuli were presented, thus obtaining food. Humans were trained with the same stimuli presented on a computer screen and had to click with a computer mouse on positive stimuli. Results showed that humans were much faster at learning to discriminate the two Greebles. In the first test, pigeons and humans had to discriminate new rotational views of the Greebles. Humans performed equally well on interpolated test views (i.e. views that lay between the training views) and extrapolated views (i.e. views outside of training range), while pigeons performed better on interpolated than on extrapolated test views. Therefore, it can be concluded that object recognition was viewpointindependent for humans and viewpoint-dependent for pigeons. In the second test, following a procedure by Aust and Huber (2006), pigeons were presented with parts of the Greebles that were not included in training and the first test to see whether they formed associations between the 2D images and the 3D objects in their aviary. They did not discriminate these parts correctly. The test was repeated with three of the test views already used in the second test but presented in different sizes. Discrimination seemed to depend on the visibility of the append

We review the cranial morphology of the Late Carboniferous terrestrial tetrapod Gephyrostegus bohemicus from the coal deposits of the Nýřany Basin in the Czech Republic. Gephyrostegus is known from several skulls ranging in length from about 25 mm to about 58 mm (holotype). The narrow skull is about twice as long as wide and shows a well-ossified quadrate and articular, but no evidence of braincase ossification. Autapomorphic features include a pustular ornamentation on some skull table bones, and a plate-like tabular process exhibiting a fine dorsal pitting. Gephyrostegus shares with Bruktererpeton fiebigi (Late Carboniferous, Germany) the presence of low, anteromedially to posterolaterally orientated sharp ridges on the posteroventral surface of the vomer. It shares with seymouriamorphs a rectangular, transverse pterygoid process and closely packed, radially arranged rows of small denticles on the palate. A phylogenetic analysis retrieves Gephyrostegidae (Gephyrostegus, Bruktererpeton) as sister group to Seymouriamorpha, although this wider clade receives low bootstrap support.

This chapter describes the distribution, population dynamics, host range and feeding preference of H. armigera and H. punctigera, the 2 most important Helicoverpa pests in Australia; host selection by Helicoverpa spp.; extent of damage on crops caused by Helicoverpa spp.; management of Helicoverpa spp. in Australia, with emphasis on the development of transgenic cotton harbouring a Bacillus thuringiensis gene, and implementation of an integrated control system incorporating INGARD (transgenic cotton); and role of the wild relatives of chickpea as sources of resistance to Helicoverpa.

Acanthostega is one of the earliest and most primitive limbed vertebrates. Its numerous fish-like features indicate a primarily aquatic lifestyle, yet cranial suture morphology suggests that its skull is more similar to those of terrestrial taxa. Here, we apply geometric morphometrics and two-dimensional finite-element analysis to the lower jaws of Acanthostega and 22 other tetrapodomorph taxa in order to quantify morphological and functional changes across the fish–tetrapod transition. The jaw of Acanthostega is similar to that of certain tetrapodomorph fish and transitional Devonian taxa both morphologically (as indicated by its proximity to those taxa in morphospace) and functionally (as indicated by the distribution of stress values and relative magnitude of bite force). Our results suggest a slow tempo of morphological and biomechanical changes in the transition from Devonian tetrapod jaws to aquatic/semi-aquatic Carboniferous tetrapod jaws. We conclude that Acanthostega retained a primitively aquatic lifestyle and did not possess cranial adaptations for terrestrial feeding.

The understanding of Earth’s biodiversity depends critically on the accurate identification and nomenclature of species. Many species were described centuries ago, and in a surprising number of cases their nomenclature or type material remain unclear or inconsistent. A prime example is provided by Elephas maximus, one of the most iconic and well-known mammalian species, described and named by Linnaeus (1758) and today designating the Asian elephant. We used morphological, ancient DNA (aDNA), and high-throughput ancient proteomic analyses to demonstrate that a widely discussed syntype specimen of E. maximus, a complete foetus preserved in ethanol, is actually an African elephant, genus Loxodonta. We further discovered that an additional E. maximus syntype, mentioned in a description by John Ray (1693) cited by Linnaeus, has been preserved as an almost complete skeleton at the Natural History Museum of the University of Florence. Having confirmed its identity as an Asian elephant through both morphological and ancient DNA analyses, we designate this specimen as the lectotype of E. maximus.

Cynodont therapsids diversified extensively after the Permo-Triassic mass extinction event, and gave rise to mammals in the Jurassic. We use an enlarged and revised dataset of discrete skeletal characters to build a new phylogeny for all main cynodont clades from the Late Permian to the Early Jurassic, and we analyse models of morphological diversification in the group. Basal taxa and epicynodonts are paraphyletic relative to eucynodonts, and the latter are divided into cynognathians and probainognathians, with tritylodonts and mammals forming sister groups. Disparity analyses reveal a heterogeneous distribution of cynodonts in a morphospace derived from cladistic characters. Pairwise morphological distances are weakly correlated with phylogenetic distances. Comparisons of disparity by groups and through time are non-significant, especially after the data are rarefied. A disparity peak occurs in the Early/Middle Triassic, after which period the mean disparity fluctuates little. Cynognathians were characterized by high evolutionary rates and high diversity early in their history, whereas probainognathian rates were low. Community structure may have been instrumental in imposing different rates on the two clades.

The tetrapod Caerorhachis bairdi, probably from the Pendleian Limestone Coal Group in the Scottish Midland Valley, is redi agnosed and redescribed, and its affinities are discussed. Caerorachis was originally interpreted as a temnospondyl amphibian, based on characters that are now regarded as primitive for tetrapods, or of uncertain polarity. Several features of Caerorhachis (e.g. gastrocentrous vertebrae, curved trunk ribs, reduced dorsal iliac blade, L-shaped tarsal intermedium) are observed in certain primitive amniotes. In particular, Caerorhachis resembles ‘anthracosaurs’, generally considered to be among the most primitive of stem-group amniotes. The phylogenetic position of Caerorhachis is considered in the light of recently published cladistic analyses of Palaeozoic tetrapods. Most analyses place Caerorhachis at the base of, or within, ‘anthra- cosaurs’. When multiple, equally parsimonious solutions are found, its ‘anthracosaur’ affinities are shown in at least some trees, and are supported by several informative and, generally, highly consistent characters. Alternative phylogenetic placements (e.g. sister taxon to temnospondyls) are usually less well corroborated. If the fundamental evolutionary split of most early tetrapods into stem-group lissamphibians (e.g. temnospondyl s) and stem-group amniotes (e.g. ‘anthracosaurs’) is accepted, then the revised interpretation of Caerorhachi s sheds light on near-ancestral conditions for Amniota.

The origin of digit-bearing tetrapods in the Middle to Late Devonian (ca. 393–360 Mya) and their subsequent invasion of land represent a classic case of a major evolutionary radiation driven by new ecological opportunities. In this and other examples, exploration of new eco-space is hypothesized to correlate with functional innovation and adaptive divergence of phenotypes. Simultaneous changes in all morphofunctional systems are rare in major evolutionary transitions and may be non-existent. Here, we focus on the mandibles of early tetrapods and their kin as a model system to test whether shifts in functional innovation were coeval with some major events in tetrapod history. To this end, we quantified mechanical variation in the mandibles of tetrapodomorphs ranging in age from Early Devonian to earliest Permian. Biomechanical disparity is stable from the Devonian to the Early Pennsylvanian, even though the origin of weight-bearing, digited limbs, and the initial phases of the colonization of land occurred during this interval. An appreciable increase in functional variation is detected in the latest Pennsylvanian and earliest Permian, when stem and crown amniotes began to explore new regions of mechanical morphospace, a pattern partly attributable to the origin of herbivory. We find no difference in the rate of functional change between tetrapodomorph “fish” and early digited tetrapods, although two independent shifts are detected among Devonian stem tetrapods more crownward than Acanthostega. Instead, the most profound shifts in evolutionary rate are nested well within the tetrapod crown and are associated with amniotes (particularly diadectomorphs and some synapsids). The substantial temporal gap between the origin of postcranial features associated with terrestriality, such as limbs with functional elbow/knee and wrist/ankle joints capable of weight-bearing, and the onset of divergence in jaw biomechanics provides a compelling example of “functional modularity” d

The bibliographic history of Martin Lister's Historiae Conchyliorum (1685-1692) is a complex one. This work, the first comprehensive work on conchology, went through several draft and partial editions. In 2012, four boxes of Listeriana were discovered in the Sackler Library at the University of Oxford. These primary sources further illuminate the techniques that Lister and his daughters utilized in constructing the Historiae, as well as the methodology they used to arrange the images of the shells for taxonomic purposes.

Adaptive radiations are central to macroevolutionary theory. Whether triggered by acquisition of new traits or ecological opportunities arising from mass extinctions, it is debated whether adaptive radiations are marked by initial expansion of taxic diversity or of morphological disparity (the range of anatomical form). If a group rediversifies following a mass extinction, it is said to have passed through a macroevolutionary bottleneck, and the loss of taxic or phylogenetic diversity may limit the amount of morphological novelty that it can subsequently generate. Anomodont therapsids, a diverse clade of Permian and Triassic herbivorous tetrapods, passed through a bottleneck during the end-Permian mass extinction. Their taxic diversity increased during the Permian, declined significantly at the Permo–Triassic boundary and rebounded during the Middle Triassic before the clade's final extinction at the end of the Triassic. By sharp contrast, disparity declined steadily during most of anomodont history. Our results highlight three main aspects of adaptive radiations: (i) diversity and disparity are generally decoupled; (ii) models of radiations following mass extinctions may differ from those triggered by other causes (e.g. trait acquisition); and (iii) the bottleneck caused by a mass extinction means that a clade can emerge lacking its original potential for generating morphological variety.

Nick Royle and Tom Pike discuss whether animals know how attractive they are. They gathered a group of females that showed clear, unambiguous preferences for males wearing red leg rings. They then showed male birds to them under four sets of experimental conditions. First, they gave the males a red leg ring, and showed them to females on the other side of a transparent partition. When the males and females could see each other, they gave each other more attention when males were wearing their attractive red leg ring-the female gave positive feedback and the male responded by courting her more. When the males wore green leg rings they got less positive feedback and spent less time courting. But when the females did not respond to the males because they could not see the amount of time the males spend courting was not related to which ring he was wearing.

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The anomalocystitid mitrate Placocystella africana (Gydo and Voorstehoek shales, upper Emsian-lower Eifelian, Bokkeveld Group, Cape Province, South Africa) is redescribed. The internal anatomy of the head, the tail morphology and part of the nervous system are reconstructed for the first time. P. africana belongs to the family Allanicytidiidae, together with four other Silurian and Devonian species from Gondwana. the monophyly of the Allanicytidiidae is supported by cladistic analysis. A second mitrate from the upper Emsian of the Bokkeveld Group, Bokkeveldia oosthuizeni gen. et sp, nov. from the Gydo Shale, is described. The distinctive ventral head skeleton of thi mitrate consists of 22 plates arranged in five transverse rows and is used as a reference to establish plate homologies in the anomalocystitids.

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A recent study proposed that incubation behaviour (i.e. type of parental care) in theropod dinosaurs can be inferred from an allometric analysis of clutch volume in extant birds. However, the study in question failed to account for factors known to affect egg and clutch size in living bird species. A new scaling analysis of avian clutch mass demonstrates that type of parental care cannot be distinguished by conventional allometry because of the confounding effects of phylogeny and hatchling maturity. Precociality of young but not paternal care in the theropod ancestors of birds is consistent with the available data.

Iguanian lizards comprise two of the most species-rich vertebrate genera on Earth (Anolis and Liolaemus). Therefore, studies with the aim of understanding their diversity and phylogenetic relationships may have major significance for ecological and evolutionary research. However, difficulties are often associated with these diverse groups. For example, adaptive radiations may lead to the evolution of conspicuous patterns of intraspecific (interpopulational) variation in response to local environmental conditions, in the absence of real speciation events. This can lead to the taxonomic recognition of new species in the absence of true reproductive isolation. In addition, although diverse taxa are appropriate models to evaluate comparatively the effects of selection on ecological and life-history traits, it is often a major challenge to gather all the available information on the distribution of these characteristics across species. This necessitates the development of synthetic works. Here we present a monographic catalogue of the diversity and phylogenetic structure of the entire South American iguanian family Liolaemidae, based on previously published studies. We also provide a complete table to summarize the distribution by country, elevational range, diet and reproductive mode of each species for which this information is available. The Liolaemidae family currently consists of a total of 229 species and subspecies belonging to the genera Ctenoblepharys, Liolaemus and Phymaturus. Remarkably, the genus Liolaemus alone comprises 209 of these taxa, consisting of 200 species, five of them polytypic, and recognized on the basis of 14 subspecies. Liolaemus species occur in Argentina, Bolivia, Brazil, Chile, Paraguay, Peru and Uruguay, representing the widest range of environments occupied by a single lizard genus. In contrast, the genus Ctenoblepharys is monotypic (Ctenoblepharys adspersa) and endemic to Peru, while 19 species of Phymaturus are distributed in Argentina

Weaving behaviour involves the repetitive lateral swaying of the head, neck, forequarters and sometimes hindquarters of the horse and is generally believed to be indicative of poor welfare. The behaviour of six known weavers was recorded three times a day for 5 days in each of three different stable designs. These were a conventional loose-box, a conventional loose-box with a 1 m2 acrylic mirror and a conventional loose-box in which there was a grilled 1 m2 side window separating the resident horse from a non-weaving conspecific in an adjacent stable. Weaving and other stereotypic behaviours were significantly higher in the unmodified stable and during the late afternoon observation period. There was no significant difference in the amount of stereotypic behaviour recorded in the two modified stables. Significant differences in the behaviour patterns and location of horses during the study suggest that activity engaging with either a visual image of a horse or a hay net is associated with a reduction in weaving and other repetitive activities in the stabled horse.

Veterinary behavioural medicine is marked by inconsistency in the naming and description of animal behavioural disorders, potentially hindering research and reducing the clinical utility of diagnoses. Sources of diagnostic inconsistency are suggested and parallels with human psychiatry are identified. This paper questions the use of categorical models as the basis for classification and reviews criticisms of categorical approaches in psychiatry. Evidence is presented to suggest an inconsistency between discrete categories and the structure of animal behavioural disorders. The potential benefits of exploring alternative models are discussed. It is concluded that it is important to move away from an approach based on advocacy and towards objective assessment of all available data. Existing diagnoses should be viewed as hypothetical constructs, with the underlying hypotheses being identified and subject to investigation. It is emphasized that global communication, empirical investigations and critical review are necessary for the development of a solid scientific basis within veterinary behavioural medicine.

Quaternary climatic fluctuations have had profound effects on the phylogeographic structure of many species. Classically, species were thought to have become isolated in peninsular refugia, but there is limited evidence that large, non-polar species survived outside traditional refugial areas. We examined the phylogeographic structure of the red fox (Vulpes vulpes), a species that shows high ecological adaptability in the western Palaearctic region. We compared mitochondrial DNA sequences (cytochrome b and control region) from 399 modern and 31 ancient individuals from across Europe. Our objective was to test whether red foxes colonised the British Isles from mainland Europe in the late Pleistocene, or whether there is evidence that they persisted in the region through the Last Glacial Maximum. We found red foxes to show a high degree of phylogeographic structuring across Europe and, consistent with palaeontological and ancient DNA evidence, confirmed via phylogenetic indicators that red foxes were persistent in areas outside peninsular refugia during the last ice age. Bayesian analyses and tests of neutrality indicated population expansion. We conclude that there is evidence that red foxes from the British Isles derived from central European populations that became isolated after the closure of the landbridge with Europe.

Metriorhynchoid crocodylians represent the pinnacle of marine specialization within Archosauria. Not only were they a major component of the Middle Jurassic–Early Cretaceous marine ecosystems, but they provide further examples that extinct crocodilians did not all resemble their modern extant relatives. Here, we use a varied toolkit of techniques, including phylogenetic reconstruction, geometric morphometrics, diversity counts, discrete character disparity analysis, and biomechanical finite-element analysis (FEA), to examine the macroevolutionary history of this clade. All analyses demonstrate that this clade became more divergent, in terms of biodiversity, form, and function, up until the Jurassic–Cretaceous boundary, after which there is no evidence for recovery or further radiations. A clear evolutionary trend towards hypercarnivory in Dakosaurus is supported by phylogenetic character optimization, morphometrics, and FEA, which also support specialized piscivory within Rhacheosaurus and Cricosaurus. Within Metriorhynchoidea, there is a consistent trend towards increasing marine specialization, with the hypermarine Cricosaurus exhibiting numerous convergences with other Mesozoic marine reptiles (e.g. loss of the deltopectoral crest and retracted external nares). In addition, biomechanics, morphometrics, and characterdisparity analyses consistently distinguish the two newly erected metriorhynchid subfamilies. This study illustrates that together with phylogeny, quantitative assessment of diversity, form, and function help elucidate the macroevolutionary pattern of fossil clades.

Inbreeding is generally rare in large, natural populations yet mate choice often appears to be random with respect to kinship. This suggests that the risks of inbreeding may be small because passive mechanisms of inbreeding avoidance, for example dispersal, are effective at lowering inbreeding risk. Previous theoretical and empirical studies have assumed that the risks of inbreeding are constant over an individual’s life span, but in the lek-breeding black grouse, inbreeding increases with female age. To determine whether inbreeding avoidance mechanisms are also age dependent, we generated four null models of random mate choice ranging from complete randomness to more biologically realistic mate choice criteria and compared these to 8 years of data on inbreeding levels at four different female age classes. We additionally tested whether mate fidelity decreased inbreeding risk. Observed inbreeding in female age classes 1, 2 or <3 were not significantly different from random, but was approximately 3.5 times higher in female age class <4. Alternative models using mate choice criteria showed no significant differences between observed and expected levels for any age class. Our results are in line with previous studies on noncooperatively breeding passerine birds, which indicate that mating is random with respect to kinship and that increased inbreeding in older females can be explained by reduced male availability caused by restrictive mate choice criteria. We also found that individuals that switched mates significantly increased relatedness to partners suggesting that mate fidelity may have evolved as an important passive inbreeding avoidance mechanism.

Additional mechanical preparation of the type material of the temnospondyl amphibian Perryella olsoni (Lower Permian, Wellington Formation, Oklahoma) highlights new cranial and postcranial features and provides additional data on previously known structures. Important new information is available for the quadrate, palatal bones and their associated dentition, parasphenoid, and appendicular skeleton. The revised diagnosis and redescription of Perryella provide the basis for a re-evaluation of its systematic affinities. A cladistic analysis of (mostly) Carboniferous and Permian temnospondyls, together with several Devonian and Carboniferous stem tetrapod outgroups, supports a single origin for temnospondyls. The sequence of branching events within temnospondyls consists of: (1) a paraphyletic Edopoidea; (2) a clade of Zatracheidae, Eryopidae, and basal Archegosauriformes; (3) a monophyletic Dvinosauria; and (4) a monophyletic Dissorophoidea. Perryella is nested within Dvinosauria in an intermediate position between Trimerorhachidae and Dvinosauroidea.

Previously described and new specimens of the anthracosaur Silvanerpeton miripedes from the Scottish Vise´an of East Kirkton yield important new data which allow us to provide a more complete reconstruction of the skull roof, palate, braincase and lower jaw. A stout sacral rib and an incompletely ossified tarsus with a subquadrangular intermedium are also recorded for the first time. A remarkably well preserved humerus in extensor view shows similarities with humeri of immature specimens of the embolomere Proterogyrinus. A new cladistic analysis, built from combining characters used in two recent matrices, places Silvanerpeton in a basal position relative to embolomeres and more derived stem amniotes. Data from Silvanerpeton inform character polarity near the base of the amniote total group. We discuss some morphofunctional implications of character changes at the root of total group amniotes, acquisition of terrestrial habits, and patterns of early disparity in this clade.

The period following the withdrawal of parental care has been highlighted as a key developmental period for juveniles. One reason for this is that juveniles cannot forage as competently as adults, potentially placing them at greater risk from environmentally-induced changes in food availability. However, no study has examined this topic. Using a long-term dataset on red foxes (Vulpes vulpes), we examined (i) dietary changes that occurred in the one-month period following the attainment of nutritional independence, (ii) diet composition in relation to climatic variation, and (iii) the effect of climatic variation on subsequent full-grown mass. Diet at nutritional independence contained increased quantities of easy-to-catch food items (earthworms and insects) when compared with pre-independence. Interannual variation in the volume of rainfall at nutritional independence was positively correlated to the proportion of earthworms in cub diet. Pre-independence cub mass and rainfall immediately following nutritional independence explained a significant proportion of variance in full-grown mass, with environmental variation affecting full-grown mass of the entire cohorts. Thus, weather-mediated availability of easyto- catch food items at a key developmental stage has lifelong implications for the development of juvenile foxes by affecting full-grown mass, which in turn appears to be an important component of individual reproductive potential.

The holotype of the brachyopoid temnospondyl Hadrokkosaurus bradyi, represented by a right lower jaw ramus, is re−ex− amined based upon new data and revision of various morphological features. Additional fragmentary jaw material re− ferred to this species is briefly described. Prominent features are a large postsymphyseal foramen that is anteriorly open, and prearticular and surangular buttresses for support of the articular. Brachyopoid characters include a long and robust postglenoid area formed by surangular and prearticular, anterior and posterior keels on at least some marginal dentary teeth, and subtriangular outline of the adductor fossa in dorsal view. Five features of the holotype ramus, long thought to be at odds with its brachyopoid or temnospondyl nature, are critically re−evaluated. A phylogenetic analysis of lower jaw characters in temnospondyls retrieves most of the clades found in more comprehensive data sets, but the statistical node support is low. Brachyopoids are monophyletic, with Hadrokkosaurus emerging as their most basal taxon.

A recent phylogenetic analysis of procolophonid parareptiles is used as the basis for a study of morphological diversity (disparity) in these amniotes. Disparity values are compared in three groups of procolophonids (a paraphyletic series of basal taxa and two monophyletic sister groups: procolophonines and leptopleuronines), two ecophenotypic assemblages (one based upon inferred diet – non high-fibre versus high-fibre species; the other based upon cranial sculpture – non horned versus horned species), and two temporal assemblages (Lower Triassic versusMiddle and Upper Triassic). The mean disparity values are comparable in the case of temporal and ecophenotypic assemblages. High-fibre species are marginally less disparate than non high-fibre species. The combined Middle and Upper Triassic species are slightly less disparate than Lower Triassic species. Finally, horned species are only slightly more disparate than non-horned species. The paraphyletic series of basal taxa and the leptopleuronines show similar disparity values, marginally higher than those for procolophonines. Phylogenetic analysis is also used to reconstruct the biogeographical history of procolophonids. Both ancestral area analysis and dispersal–vicariance analysis show that South Africa was the most likely ancestral area for procolophonids as a whole. North China – either as a single area or in combination with Russia or South Africa – was the most likely ancestral area for the leptopleuronine–procolophonine clade.

In an attempt to investigate differences between the most widely discussed hypotheses of early tetrapod relationships, we assembled a new data matrix including 90 taxa coded for 319 cranial and postcranial characters. We have incorporated, where possible, original observations of numerous taxa spread throughout the major tetrapod clades. A stem-based ( total-group) definition of Tetrapoda is preferred over apomorphy- and node-based (crown-group) definitions. This definition is operational, since it is based on a formal character analysis. A PAUP* search using a recently implemented version of the parsimony ratchet method yields 64 shortest trees. Differences between these trees concern: (1) the internal relationships of aı¨stopods, the three selected species of which form a trichotomy; (2) the internal relationships of embolomeres, with Archeria crassidisca and Pholiderpeton scutigerum collapsed in a trichotomy with a clade formed by Anthracosaurus russelli and Pholiderpeton attheyi ; (3) the internal relationships of derived dissorophoids, with four amphibamid species forming an unresolved node with a clade consisting of micromelerpetontids and branchiosaurids and a clade consisting of albanerpetontids plus basal crown-group lissamphibians ; (4) the position of albenerpetontids and Eocaecilia micropoda, which form an unresolved node with a trichotomy subtending Karaurus sharovi, Valdotriton gracilis and Triadobatrachus massinoti ; (5) the branching pattern of derived diplocaulid nectrideans, with Batrachiderpeton reticulatum and Diceratosaurus brevirostris collapsed in a trichotomy with a clade formed by Diplocaulus magnicornis and Diploceraspis burkei. The results of the original parsimony run – as well as those retrieved from several other treatments of the data set ( e.g. exclusion of postcranial and lower jaw data ; character reweighting ; reverse weighting ) – indicate a deep split of early tetrapods between lissamphibian- and amniote-related taxa. Colosteids

Priapulids and their extinct relatives, the archaeopriapulids and palaeoscolecids, are vermiform, carnivorous ecdysozoans with an armoured, extensible proboscis. These worms were an important component of marine communities during the Palaeozoic, but were especially abundant and diverse in the Cambrian. Today, they comprise just seven genera in four families. Priapulids were among the first groups used to test hypotheses concerning the morphological disparity of Cambrian fossils relative to the extant fauna. A previous study sampled at the generic level, concluding that Cambrian genera embodied marginally less morphological diversity than their extant counterparts. Here, we sample predominantly at the species level and include numerous fossils and some extant forms described in the last fifteen years. Empirical morphospaces for priapulids, archaeopriapulids and palaeoscolecids are relatively insensitive to changes in the taxon or character sample: their overall form has altered little, despite the markedly improved sampling. Cambrian and post-Cambrian genera occupy adjacent rather than broadly overlapping regions of these spaces, and Cambrian species still show lower morphological disparity than their post-Cambrian counterparts. Crucially, the significance of this difference has increased with improved taxon sampling over research time. In contrast with empirical morphospaces, the phylogeny of priapulids, archaeopriapulids and palaeoscolecids derived from morphological characters is extremely sensitive to details of taxon sampling and the manner in which characters are weighted. However, the extant Priapulidae and Halicryptidae invariably resolve as sister families, with this entire clade subsequently being sister group to the Maccabeidae. In our most inclusive trees, the extant Tubiluchidae are separated from these other living taxa by a number of small, intervening fossil clades.

Temnospondyls—a major component of Permian and Carboniferous terrestrial ecosystems—display great diversity in skull shapes and proportions. To quantify and interpret this diversity, we conducted a geometric morphometric analysis using 45 landmarks on the dorsal skull surface of 90 species with well-represented cranial material. Results show a correlation between morphospace occupation and phylogenetic proximity of taxa for trees in which dvinosaurs and dissorophoids are sister groups and join an edopoid-–eryopoid–basal archegosauriform clade. Most large groups of Carboniferous and Permian temnospondyls occupy specific areas of morphospace. Nearest-neighbor analyses reveal significantly greater taxon clustering than expected under either uniform or Gaussian null models. Size correlates strongly with shape across the whole data set, highlighting the association of some features (short, broad snout; large orbits) with small size. A significant relationship between size and shape is not observed in clades such as branchiosaurids, dvinosaurs, and olsoniforms (the clade encompassing trematopids plus dissorophids). This suggests that evolutionary allometry patterns vary across temnospondyls. In the case of branchiosaurids, this pattern may be explained by the similar sizes and relatively conservative morphologies of the constituent species. Distance-based disparity measures indicate that edopoids, eryopoids, and basal archegosauriforms make the largest contributions to total disparity (reflecting the peripheral locations of some of the constituent taxa in morphospace), whereas amphibamids, dissorophids, and trematopids make the smallest contributions. Disparity correlates strongly with diversity within groups, suggesting that skull shape was not subject to character state exhaustion (decrease or cessation in the acquisition of novel morphological conditions during clade evolution). The Kasimovian, Roadian, Wordian, and Changhsingian are time intervals of high disparity de

It is important to understand the quality of the fossil record of early tetrapods (Tetrapoda, minus Lissamphibia and Amniota) because of their key role in the transition of vertebrates from water to land, their dominance of terrestrial faunas for over 100 million years of the late Palaeozoic and earlyMesozoic, and their variable fates during the end−Permian mass extinction. The first description of an early tetrapod dates back to 1824, and since then discoveries have occurred at a rather irregular pace, with peaks and troughs corresponding to some of the vicissitudes of human history through the past two centuries. As expected, the record is dominated by the well−sampled sedimentary basins of Europe and North America, but finds from other continents are increasing rapidly. Comparisons of snapshots of knowledge in 1900, 1950, and 2000 show that discovery of new species has changed the shape of the species−level diversification curve, contrary to earlier studies of family−level taxa. There is, however, little evidence that taxon counts relate to research effort (as counted by numbers of publications), and there are no biasing effects associated with differential study of different time intervals through the late Palaeozoic and Mesozoic. In fact, levels of effort are apparently not related to geological time, with no evidence that workers have spent more time on more recent parts of the record. In particular, the end−Permian mass extinction was investigated to determine whether diversity changes through that interval might reflect worker effort: it turns out that most records of early tetrapod taxa (when corrected for duration of geological series) occur in the Lower Triassic.

Extant amphibians consist of Salientia (frogs), Caudata (salamanders), and Gymnophiona (caecilians). The mutual relationships of these groups are controversial, with either Batrachia (Salientia + Caudata) or Procera (Gymnophiona + Caudata) as emerging clades in recent molecular and morphological analyses. The monophyly of amphibians as a whole is supported by independent data, but their origins and affinities with early tetrapods are debated. A new cladistic analysis of early tetrapods retrieves Temnospondyli (the most species-rich group of early tetrapods) as the closest relatives of crown group amphibians. One temnospondyl group, the Dissorophoidea, forms a series of consecutive outgroups to crown amphibians. In particular, the Lower Permian amphibamid Doleserpeton is the most derived plesion on the amphibian stem. The Albanerpetontidae, a group of salamander-like tetrapods ranging from the Jurassic to the Pliocene, are placed as stem Gymnophiona. The shortest trees support the Batrachia hypothesis. However, the Procera hypothesis is not a significantly worse fit for the whole character set. Exhaustive treatment of characters and taxa is the most appropriate way to disentangle contrasting phylogenetic signals in large matrices. Tests of different crown topologies show that placement of amphibians within lepospondyls (e.g. as sister taxon to Lysorophia) is not a significantly worse fit for the whole character set than a close temnospondyl-lissamphibian relationship. However, the latter phylogenetic hypothesis best captures the most coherent assembly of derived lissamphibian apomorphies.

Ichthyosaurs were important marine predators in the Early Jurassic, and an abundant and diverse component of Mesozoic marine ecosystems. Despite their ecological importance, however, the Early Jurassic species represent a reduced remnant of their former significance in the Triassic. Ichthyosaurs passed through an evolutionary bottleneck at, or close to, the Triassic-Jurassic boundary, which reduced their diversity to as few as three or four lineages. Diversity bounced back to some extent in the aftermath of the end-Triassic mass extinction, but disparity remained at less than one-tenth of pre-extinction levels, and never recovered. The group remained at low diversity and disparity for its final 100 Myr. The end-Triassic mass extinction had a previously unsuspected profound effect in resetting the evolution of apex marine predators of the Mesozoic.

A maximum parsimony analysis of a data matrix including 42 temnospondyl species and 14 outgroup species coded for 246 characters results in eight shortest trees. Temnospondyls emerge as two distinct radiations: one includes edopoids and eryopoids-basal archegosauriforms; the other includes dissorophoids and dvinosaurs. Both receive low statistical support. The branching pattern of dissorophoids remains elusive and is obscured by recurrent homoplasies. The two Micropholis morphs appear as sister taxa near the base of the dissorophoid clade. Perryella is either placed as sister taxon to dvinosaurs or nested within them. Temnospondyls such as the genera Balanerpeton and Dendrerpeton occur close to the node that subtends the dvinosaur–dissorophoid separation. Disparity analysis using Principal Coordinate Analysis of both Manhattan and Euclidean intertaxon distance matrices reveals that all major temnospondyl groups are widely separated in morphospace. Levels of disparity are comparable in the various clades examined, but the dvinosaurs tend to be morphologically more diverse than other groups when variance-derived metrics are used. For range-derived metrics, dissorophoids are more disparate than other groups. Overall disparity is similar in edopopids and eryopoids-basal archegosauriforms, as is in dissorophoids and dvinosaurs. However, the latter two groups are invariably slightly more disparate than the former two.